Visual resolution,contrast sensitivity,and the cortical magnification factor

Summary This study shows that photopic contrast sensitivity and resolution can be predicted by means of simple functions derived by using the cortical magnification factor M as a scale factor of mapping from the visual field into the striate cortex. We measured the minimum contrast required for discriminating the direction of movement or orientation of sinusoidal gratings, or for detecting them in central and peripheral vision. No qualitative differences were found between central and peripheral vision, and almost all quantitative differences observed could be removed by means of a size compensation derived from M.The results indicated specificly that (1) visual patterns can be made equally visible if they are scaled so that their calculated cortical representations become equivalent; (2) contrast sensitivity follows the same power function of the cortical area stimulated by a grating at any eccentricity; (3) area and squared spatial frequency are reciprocally related as determinants of contrast sensitivity; and (4) acuity and resolution are directly proportional to M, and the minimum angle of resolution is directly proportional to M^-1.The power law of spatial summation expressed in (2) and (3) suggests the existence of a central integrator that pools the activity of cortical neurons. This summation mechanism makes the number of potentially activated visual cells the most important determinant of visibility and contrast sensitivity. The functional homogeneity of image processing across the visual field observed here agrees with the assumed anatomical and physiological uniformity of the visual cortex.     

Vision influences on tactile discrimination of grating orientation

Studies have revealed a number of instances of cross-modal spatial interactions between vision and touch. However, few studies have addressed the possible role of cross-modal interactions in the identification of objects. This study assessed whether tactile pattern perception is affected by adding a concurrent visual stimulus. Participants were required to discriminate the orientation of tactile grating patterns, which were presented in synchrony or asynchrony with a task-irrelevant visual grating pattern that had the same or different orientation as the tactile pattern. The results demonstrated that accuracy in discriminating grating orientation was reduced when the tactile and visual gratings differed in orientation, but only when the two modalities were presented concurrently. The findings are discussed in terms of multisensory integration for processing of shape information.     

BOLD fMRI response of early visual areas to perceived contrast in human amblyopia

In this study, we used a temporal two-alternative forced choice psychophysical procedure to measure the observer's perception of a 22% physical contrast grating for each eye as a function of spatial frequency in four subjects with unilateral amblyopia and in six subjects with normal vision. Contrast thresholds were also measured using a standard staircase method. Additionally, blood-oxygenation-level-dependent (BOLD) functional magnetic resonance imaging (fMRI) was used to measure the neuronal response within early visual cortical areas to monocular presentations of the same 22% physical contrast gratings as a function of spatial frequency. For all six subjects with normal vision and for three subjects with amblyopia, the psychophysically measured perception of 22% contrast as a function of spatial frequency was the same for both eyes. Threshold contrast, however, was elevated for the amblyopic eye for all subjects, as expected. The magnitude of the fMRI response to 22% physical contrast within "activated" voxels was the same for each eye as a function of spatial frequency, regardless of the presence of amblyopia. However, there were always fewer "activated" fMRI voxels during amblyopic stimulation than during normal eye stimulation. These results are consistent with the hypotheses that contrast thresholds are elevated in amblyopia because fewer neurons are responsive during amblyopic stimulation, and that the average firing rate of the responsive neurons, which reflects the perception of contrast, is unaffected in amblyopia.     

Contrast sensitivity in humans with abnormal visual experience.

1. Grating contrast sensitivities have been determined over a range of spatial frequencies for a normal subject and for subjects who are visually biased in that they have a lower resolution capacity for targets of specific orientations. The bias si only found in astigmatic subjects and the grating orientation yielding poorest acuity coincides with the most defocused astigmatic meridian. However this resolution anisotropy remains when optical factors are accounted for. 2. For the normal subject, high and low frequency attenuation is found and a typical reduction in contrast sensitivity is exhibited for oblique target orientations. 3. The biased subjects, called meridional amblyopes because they have reduced acuity for a given grating orientation, show markedly abnormal contrast sensitivity functions. Their cut-off spatial frequencies are different for various target orientations and this difference applies also to contrast sensitivity over nearly the entire spatial frequency range tested (0-5-16 cycles/deg). The differences are of about the same magnitude for most frequencies and they are found in all types of meridional amblyopes. 4. Optical explanations of these differences are ruled out by laser-interference fringe tests and by varying effective pupil size. 5. Theoretical effects of defocus have been calculated to compare predicted visual deprivation with performance. Results indicate that reduced contrast sensitivity functions can be equivalent to a small defocus effect. 6. To examine the results in the spatial domain, inverse Fourier transforms of representative contrast sensitivity functions have been computed. The optical portion of the resulting spatial weighting functions has been parcelled out to obtain neural spatial weighting functions.     

Contrast-dependent changes in spatial frequency tuning of macaque V1 neurons: effects of a changing receptive field size

Previous studies on single neurons in primary visual cortex have reported that selectivity for orientation and spatial frequency tuning do not change with stimulus contrast. The prevailing hypothesis is that contrast scales the response magnitude but does not differentially affect particular stimuli. Models where responses are normalized over contrast to maintain constant tuning for parameters such as orientation and spatial frequency have been proposed to explain these results. However, our results indicate that a fundamental property of receptive field organization, spatial summation, is not contrast invariant. We examined the spatial frequency tuning of cells that show contrast-dependent changes in spatial summation and have found that spatial frequency selectivity also depends on stimulus contrast. These results indicate that contrast changes in the spatial frequency tuning curves result from spatial reorganization of the receptive field.     

Visual evoked responses in humans with abnormal visual experience.

1. The visual evoked response to a grating target of varying spatial frequency was examined for normal subjects and for subjects with meridional amblyopia. This condition, reduced visual resolution for specific target orientations, is associated with, and thought to result from, marked ocular astigmatism. 2. For normal subjects, the general relation between spatial frequency and the evoked response is similar to psychophysical contrast sensitivity data. Evoked response amplitudes to oblique gratings are typically reduced and this is analogous to the lower acuity for oblique compared to horizontal and vertical detail. 3. In addition to the oblique effect, the magnitude of the evoked response for meridional amblyopes depends upon grating orientation over most of the spatial frequency range tested (0-5-16 cycles/deg). The lowest evoked amplitude is found when stimulus grating orientation matches that for which acuity is reduced. 4. The evoked potentials spatial frequency response functions are qualitatively similar to contrast sensitivity functions determined with the same abnormal subjects. 5. From these results, it may be concluded that the physiological locus of meridional amblyopia is confined primarily to structures at or prior to the site of evoked potential generation.     

From the heart to the mind's eye: cardiac vagal tone is related to visual perception of fearful faces at high spatial frequency

The neurovisceral integration model (Thayer and Lane, 2000) proposes that vagally mediated heart rate variability (HRV)--an index of cardiac vagal tone--is associated with autonomic flexibility and emotional self-regulation. Two experiments examined the relationship between vagally mediated HRV and visual perception of affectively significant stimuli at different spatial frequencies. In Experiment 1, HRV was positively correlated with superior performance discriminating the emotion of affectively significant (i.e., fearful) faces at high spatial frequency (HSF). In Experiment 2, processing goals moderated the relationship between HRV and successful discrimination of HSF fearful faces. In contrast to Experiment 1, discriminating the expressiveness of HSF fearful faces was not correlated with HRV. The current research suggests that HRV is positively associated with superior visual discrimination of affectively significant stimuli at high spatial frequency, and this relationship may be sensitive to the top-down influence of different processing goals.     

Spatial resolution for feature binding is impaired in peripheral and amblyopic vision

We measured spatial resolution for discriminating targets that differed from nearby distractors in either color or orientation or their conjunction. In the fovea of normal human observers, whenever both attributes are big enough to be individually visible, their conjunction is also visible. In the periphery, the two attributes may be visible, but their conjunction may be invisible. We found a similar impairment in resolving conjunctions for the fovea of deprived eyes of humans with abnormal visual development (amblyopia). These results are quantitatively explained by a model of primary visual cortex (V1) in which orientation and color maps are imperfectly co-registered topographically. Our results in persons with amblyopia indicate that the ability of the fovea to compensate for this poor co-registration is consolidated by visual experience during postnatal development.     

Binocular deficits associated with early alternating monocular defocus. I. Behavioral observations

To study the binocular vision deficits associated with anisometropia, monkeys were reared with alternating monocular defocus, which allowed monocular mechanisms to develop normally while binocular mechanisms were selectively compromised. A defocusing contact lens of -1.5 D, -3 D, or -6 D was worn on alternate eyes on successive days (n = 3 per lens power) from 3 wk to 9 mo of age. The control subjects were two normally reared monkeys and two human observers. Functional binocular vision was assessed through behavioral measurements of stereoscopic depth discrimination thresholds as a function of spatial frequency. To characterize the extent of the deficits in disparity processing at a given spatial frequency, the contrast required to support stereopsis was determined for a range of disparities that exceeded the subjects' measured stereoacuity. The lens-reared monkeys showed spatial-frequency-selective deficits in stereopsis that depended on the magnitude of the simulated anisometropia experienced during the rearing period. For a given spatial frequency, the treated monkeys generally required higher than normal contrasts to support stereopsis even for large disparities. Moreover, a given increase in contrast produced smaller than normal improvements in stereo discrimination in our treated subjects, which suggests that in addition to deficits in contrast sensitivity, disparity-sensitive mechanisms exhibited low contrast gains. The spatial-frequency selective nature of the binocular deficits produced by the imposed anisometropia indicate that disparity processing mechanisms are normally spatial-frequency selective and that mechanisms tuned to different spatial frequencies can be differentially affected by abnormal binocular visual experience.     

Visual orientation and spatial frequency discrimination: a comparison of single neurons and behavior

Neurons in the visual cortex respond selectively to stimulus orientation and spatial frequency. Changes in response amplitudes of these neurons could be the neurophysiological basis of orientation and spatial frequency discrimination. We have estimated the minimum differences in stimulus orientation and spatial frequency that can produce reliable changes in the responses of individual neurons in cat visual cortex. We compare these values with orientation and spatial frequency discrimination thresholds determined behaviorally. Slopes of the tuning functions and response variability determine the minimum orientation and spatial frequency differences that can elicit a reliable response change. These minimum values were obtained from single cells using receiver operating characteristic (ROC) analysis. The average minimum orientation and spatial frequency differences that could be signaled reliably by cells from our sample were 6.4 degrees (n = 22) and 21.3% (n = 18), respectively. These values are approximately 0.20 of the average full tuning width at one-half height of the cells. Although these average values are well above the behaviorally determined thresholds, the most selective cells signaled orientation and frequency differences of 1.84 degrees and 5.25%, respectively. These values are of the same order of magnitude as the behavioral thresholds. We show that, because of slow fluctuations in a cell's responsivity, ROC analysis overestimates response variability. We estimate that these slow response fluctuations elevated our estimates of single cell "thresholds" by, on average, 30%. Our data point to an approximate correspondence between orientation and spatial frequency discrimination "thresholds" determined behaviorally and those estimated from the most selective single cortical cells. Interpretation of this quantitative correspondence is considered in the discussion.     

The effects of contrast on visual orientation and spatial frequency discrimination: a comparison of single cells and behavior

We have compared the effects of contrast on human psychophysical orientation and spatial frequency discrimination thresholds and on the responses of individual neurons in the cat's striate cortex. Contrast has similar effects on orientation and spatial frequency discrimination: as contrast is increased above detection threshold, orientation and spatial frequency discrimination performance improves but reaches maximum levels at quite low contrasts. Further increases in contrast produce no further improvements in discrimination. We measured the effects of contrast on response amplitude, orientation and spatial frequency selectivity, and response variance of neurons in the cat's striate cortex. Orientation and spatial frequency selectivity vary little with contrast. Also, the ratio of response variance to response mean is unaffected by contrast. Although, in many cells, response amplitude increases approximately linearly with log contrast over most of the visible range, some cells show complete or partial saturation of response amplitude at medium contrasts. Therefore, some cells show a clear increase in slope of the orientation and spatial frequency tuning functions with increasing contrast, whereas in others the slopes reach maximum values at medium contrasts. Using receiver operating characteristic analysis, we estimated the minimum orientation and spatial frequency differences that can be signaled reliably as a response change by an individual cell. This analysis shows that, on average, the discrimination of orientation or spatial frequency improves with contrast at low contrasts more than at higher contrasts. Using the optimal stimulus for each cell, we estimated the contrast threshold of 48 neurons. Most cells had contrast thresholds below 5%. Thresholds were only slightly higher for nonoptimal stimuli. Therefore, increasing the contrast of sinusoidal gratings above approximately 10% will not produce large increases in the number of responding cells. The observed effects of contrast on the response characteristics of nonsaturating cortical cells do not appear consistent with the psychophysical results. Cells that reach their maximum response at low-to-medium contrasts may account for the contrast independence of psychophysical orientation and spatial frequency discrimination thresholds at medium and high contrasts.     

Visual acuity in the blue cone monochromat

1. To isolate the blue cones of the normal eye, blue sine-wave gratings were superimposed on a bright yellow background. Threshold contrasts for resolution of the gratings were then determined.2. Under these conditions visual acuity for high contrast gratings on four normal subjects was on the average 0.31 min(-1). This is about a factor of 6 lower than grating acuity under optimal conditions.3. The contrast sensitivity of two subjects who lack the normal red and green receptor mechanisms was measured using blue sinusoidally modulated gratings. Visual acuity was found to be greatly reduced from normal. The low acuity of these individuals is due to both a reduction in contrast sensitivity and a reduction in resolution.4. The spatial resolving characteristics of these subjects resembles the vision of the normal eye under conditions which isolate the blue sensitive mechanisms.5. The vision of the cone monochromat differs significantly from that of the more typical rod monochromat. The rod monochromat has even lower acuity than the blue cone monochromat for high contrast gratings but has much greater contrast sensitivity for gratings of low spatial frequency.     

Delayed discrimination of spatial frequency for gratings of different orientation: behavioral and fMRI evidence for low-level perceptual memory stores in early visual cortex

The concept of perceptual memory refers to the neural and cognitive processes underlying the storage of specific stimulus features such as spatial frequency, orientation, shape, contrast, and color. Psychophysical studies of perceptual memory indicate that observers can retain visual information about the spatial frequency of Gabor patterns independent of the orientation with which they are presented. Compared to discrimination of gratings with the same orientation, reaction times to orthogonally oriented gratings, however, increase suggesting additional processing. Using event-related fMRI we examined the pattern of neural activation evoked when subjects discriminated the spatial frequency of Gabors presented with the same or orthogonal orientation. Blood-oxygen level dependent BOLD fMRI revealed significantly elevated bilateral activity in visual areas (V1, V2) when the gratings to be compared had an orthogonal orientation, compared to when they had the same orientation. These findings suggest that a change in an irrelevant stimulus dimension requires additional processing in primary and secondary visual areas. The finding that the task-irrelevant stimulus property (orientation) had no significant effect on the prefrontal and intraparietal cortex supports a model of working memory in which discrimination and retention of basic stimulus dimensions is based on low-level perceptual memory stores that are located at an early stage in the visual process. Our findings suggest that accessing different stores requires time and has higher metabolic costs. Supported by: BMBF Project “Visuospatial Cognition” and Norwegian Research Council.     

Multisensory enhancement of localization under conditions of induced myopia

Enhanced behavioral performance mediated by multisensory stimuli has been shown using a variety of measures, including response times, orientation behaviors, and even simple stimulus detection. However, there has been little evidence for a multisensory-mediated improvement in stimulus localization. We suggest that this lack of effect may be a result of the high acuity of the visual system. To examine whether normal visual acuity may be masking any potential multisensory benefit for stimulus localization, we examined the ability of human subjects to localize visual, auditory and combined visual-auditory targets under conditions of normal and degraded vision. Under conditions of normal vision, localization precision (i.e., variability) was equivalent for visual and multisensory targets, and was significantly worse for auditory targets. In contrast, under conditions of induced myopia, visual localization performance was degraded by an average of 25%, while auditory localization performance was unaffected. However, during induced myopia, multisensory (i.e., visual-auditory) localization performance was significantly improved relative to visual performance. These results show a multisensory-mediated enhancement in human localization ability, and illustrate the cross-modal benefits that can be obtained when spatial information in one sense is compromised or ambiguous.     

Modification of the kitten's visual cortex by exposure to spatially periodic patterns

Kittens were dark-reared except for exposure to three types of spatially periodic, vertically striped pattern: 1. single, widely spaced black bars; 2. wide areas of regular vertical grating separated by large blank patches; 3. a uniform, continuous grating with a spatial frequency of 0.5 c/deg. In each case there was a bias towards vertical in the distribution of preferred orientations of cells recorded in the visual cortex. The contrast sensitivity of individual neurones for gratings of different spatial frequencies was analysed quantitatively. In kittens exposed to a uniform grating of 0.5 c/deg, many cells were maximally sensitive close to 0.5 c/deg, as they are in normal cats. The occipital potential evoked by vertical gratings higher in frequency than 0.3 c/deg was consistently greater in amplitude than that for horizontal, and a vertical grating of 0.5 c/deg produced the maximum activity. These results are compared with those of Maffei and Fiorentini (1974); the differences between our results and theirs may be attributable to the degree of variability in spatial frequency and orientation during rearing, and to the duration of exposure.     

Cortical maps of separable tuning properties predict population responses to complex visual stimuli

In the earliest cortical stages of visual processing, a scene is represented in different functional domains selective for specific features. Maps of orientation and spatial frequency preference have been described in the primary visual cortex using simple sinusoidal grating stimuli. However, recent imaging experiments suggest that the maps of these two spatial parameters are not sufficient to describe patterns of activity in different orientation domains generated in response to complex, moving stimuli. A model of cortical organization is presented in which cortical temporal frequency tuning is superimposed on the maps of orientation and spatial frequency tuning. The maps of these three tuning properties are sufficient to describe the activity in orientation domains that have been measured in response to drifting complex images. The model also makes specific predictions about how moving images are represented in different spatial frequency domains. These results suggest that the tangential organization of primary visual cortex can be described by a set of maps of separable neuronal receptive field features including maps of orientation, spatial frequency, and temporal frequency tuning properties.     

Contrast constancy: deblurring in human vision by spatial frequency channels.

The perception of contrast was measured in humans by a technique of subjective contrast-matching, and was compared with contrast sensitivity as defined by threshold measures. 2. Contrast-matching between different spatial frequencies was performed correctly (especially at frequencies above 5 c/deg) despite the attenuation by optical and neural factors which cause large differences in contrast thresholds. 3. Contrast-matching between single lines of different widths was also veridical, and was not limited by the spatial integration (Ricco's Law) present at threshold. Adaptation to gratings altered the appearance of lines, and this could be best understood in Fourier terms. 4. The generality of these results was shown by matching the contrast of pictures which had been filtered so that each contained a one octave band of spatial frequencies. 5. Within the limits imposed by threshold and resolution, contrast-matching was largely independent of luminance and position on the retina. 6. Six out of eleven astigmatic observers showed considerable suprathreshold compensation for their orientation-specific neural deficit in contrast sensitivity. 7. These results define a new property of vision: contrast constancy. It is argued that spatial frequency channels in the visual cortex are organized to compensate for earlier attenuation. This achieves a dramatic 'deblurring' of the image, and optimizes the clarity of vision.     

Intermodal spatial attention differs between vision and audition: an event-related potential analysis

Subjects were required to attend to a combination of stimulus modality (vision or audition) and location (left or right). Intermodal attention was measured by comparing event-related potentials (ERPs) to visual and auditory stimuli when the modality was relevant or irrelevant, while intramodal (spatial) attention was measured by comparing ERPs to visual and auditory stimuli presented at relevant and irrelevant spatial locations. Intramodal spatial attention was expressed differently in visual and auditory ERPs. When vision was relevant, spatial attention showed a contralateral enhancement of posterior N1 and P2 components and enhancement of parietal P3. When audition was relevant, spatial attention showed a biphasic fronto-central negativity, starting after around 100 ms. The same effects were also present in ERPs to stimuli that were presented in the irrelevant modality. Thus, spatial attention was not completely modality specific. Intermodal attention effects were also expressed differently in vision and audition. Taken together, the obtained ERP patterns of the present study show that stimulus attributes such as modality and location are processed differently in vision and audition.     

Effects of early discordant binocular vision on the postnatal development of parvocellular neurons in the monkey lateral geniculate nucleus

 The effects of early discordant binocular vision on the fidelity of signal transfer in parvocellular neurons of the lateral geniculate nucleus (LGN) were investigated in rhesus monkeys reared with ocular misalignment (strabismus). Unilateral convergent strabismus (esotropia) was surgically induced in four infant monkeys between 20 and 30 days of age and the animals were reared in a normally lighted environment until they were adults. Extracellular microelectrode recordings were made in individual units of anesthetized and paralyzed subjects. Drifting sinusoidal gratings were used as visual stimuli. Within-unit comparisons of the LGN action potentials (LGN output) and S potentials (retinal input) were performed to determine the accuracy of signal transfer in the LGN. Contrary to the previous findings in the cat LGN, the signal transfer characteristics of parvocellular units in strabismic monkeys were normal regardless of stimulus spatial frequency, temporal frequency, or contrast. The differences between cats and monkeys in LGN circuitry and the relative maturity of the central visual pathway at the onset of strabismus may have contributed to the apparent species differences in the functional development of the LGN. Received: 26 March 1997 / Accepted: 18 July 1997